{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,14]],"date-time":"2026-08-14T05:29:59Z","timestamp":1786685399932,"version":"build-2736575974"},"reference-count":38,"publisher":"Springer Science and Business Media LLC","issue":"8","license":[{"start":{"date-parts":[[2026,7,23]],"date-time":"2026-07-23T00:00:00Z","timestamp":1784764800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2026,7,23]],"date-time":"2026-07-23T00:00:00Z","timestamp":1784764800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"name":"Fundaci\u00f3 Instituci\u00f3 dels Centres de Recerca de Catalunya"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Exp Brain Res"],"published-print":{"date-parts":[[2026,8]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Precise coordination between neurons and cortical areas is crucial for sensory perception, yet in vision, continuous fixational eye movements constantly shift retinal input. Traditionally considered random, these drifts modulate neural activity and may challenge the temporal precision needed for perceptual integration, particularly given their partial interocular independence. Recent evidence suggests that ocular drifts are not purely stochastic and play an active role in visual processing. Here, we investigated the impact of fixational eye movements on visual perception using a figure\u2013ground segmentation task. Participants maintained central fixation while viewing textures composed of oriented line segments and reported the presence or absence of a centrally embedded figure. We simultaneously recorded binocular drift and microsaccades, analyzing interocular coordination through 2D cross-correlograms of angular velocities. We found that figure-presents trials exhibited narrow peaks in interocular correlation, while figure-absent trials showed broader, less synchronized drift patterns. Microsaccades were observed to facilitate drift alignment across the eyes. The decrease in microsaccade rate in figure-present trials may contribute to the change in drift synchrony, but it is not required for it. Changes in eye synchrony are also observed under monocular viewing conditions, whereas variations in accommodation and pupil size appear to be unrelated. These results identify two distinct ocular drift modes associated with different perceptual outcomes. Our findings suggest that ocular drift operates under closed-loop control and contributes to effective visual processing.<\/jats:p>","DOI":"10.1007\/s00221-026-07320-y","type":"journal-article","created":{"date-parts":[[2026,7,23]],"date-time":"2026-07-23T08:26:25Z","timestamp":1784795185000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Two distinct modes of ocular drift observed during figure-ground perception"],"prefix":"10.1007","volume":"244","author":[{"given":"August","family":"Romeo","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Maria Sol\u00e9","family":"Puig","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hans","family":"Sup\u00e8r","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,7,23]]},"reference":[{"key":"7320_CR2","doi-asserted-by":"publisher","first-page":"815","DOI":"10.1038\/383815a0","volume":"383","author":"JM Alonso","year":"1996","unstructured":"Alonso JM, Usrey WM, Reid RC (1996) Precisely correlated firing in cells of the lateral geniculate nucleus. Nature 383:815\u2013819","journal-title":"Nature"},{"key":"7320_CR3","doi-asserted-by":"publisher","first-page":"1697","DOI":"10.1038\/s41467-022-29201-y","volume":"13","author":"N Ben-Shushan","year":"2022","unstructured":"Ben-Shushan N, Shaham N, Joshua M, Burak Y (2022) Fixational drift is driven by diffusive dynamics in central neural circuitry. Nat Commun 13:1697","journal-title":"Nat Commun"},{"key":"7320_CR5","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0185180","author":"NR Bowers","year":"2017","unstructured":"Bowers NR, Poletti M (2017) Microsaccades during reading. PLoS ONE. https:\/\/doi.org\/10.1371\/journal.pone.0185180","journal-title":"PLoS ONE"},{"key":"7320_CR6","doi-asserted-by":"publisher","DOI":"10.1093\/acprof:oso\/9780195301069.001.0001","volume-title":"Rhythms of the brain","author":"G Buzs\u00e1ki","year":"2006","unstructured":"Buzs\u00e1ki G (2006) Rhythms of the brain. Oxford Univ. Press"},{"key":"7320_CR7","doi-asserted-by":"publisher","first-page":"e2200256119","DOI":"10.1073\/pnas.2200256119","volume":"119","author":"AM Clark","year":"2022","unstructured":"Clark AM, Intoy J, Rucci M, Poletti M (2022) Eye drift during fixation predicts visual acuity. Proc Natl Acad Sci U S A 119:e2200256119. https:\/\/doi.org\/10.1073\/pnas.2200256119","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"48","key":"7320_CR8","doi-asserted-by":"publisher","first-page":"e0594242024","DOI":"10.1523\/JNEUROSCI.0594-24.2024","volume":"44","author":"AM Clark","year":"2024","unstructured":"Clark AM, Huynh A, Poletti M (2024) Oculomotor contributions to foveal crowding. J Neurosci 44(48):e0594242024. https:\/\/doi.org\/10.1523\/JNEUROSCI.0594-24.2024","journal-title":"J Neurosci"},{"key":"7320_CR9","doi-asserted-by":"publisher","first-page":"137","DOI":"10.1016\/S0028-3932(98)00088-8","volume":"37","author":"E Corthout","year":"1999","unstructured":"Corthout E, Uttl B, Ziemann U, Cowey A, Hallett M (1999) Two periods of processing in the (circum)striate visual cortex as revealed by transcranial magnetic stimulation. Neuropsychologia 37:137\u2013145","journal-title":"Neuropsychologia"},{"key":"7320_CR10","doi-asserted-by":"publisher","first-page":"121","DOI":"10.1007\/BF00202899","volume":"60","author":"R Eckhorn","year":"1988","unstructured":"Eckhorn R, Bauer R, Jordan W, Brosch M, Kruse W, Munk M, Reitboeck H (1988) Coherent oscillations: a mechanism of feature linkingin the visual cortex? Multiple electrode and correlation analysesin the cat. Biol Cybern 60:121\u2013130","journal-title":"Biol Cybern"},{"key":"7320_CR11","doi-asserted-by":"publisher","first-page":"56","DOI":"10.1073\/pnas.1102730108","volume":"2","author":"R Engbert","year":"2011","unstructured":"Engbert R, Mergenthaler K (2011) An integrated model of fixational eye movements and microsaccades. PNAS 2:56. https:\/\/doi.org\/10.1073\/pnas.1102730108","journal-title":"PNAS"},{"key":"7320_CR12","doi-asserted-by":"publisher","first-page":"1035","DOI":"10.1016\/S0042-6989(03)00084-1","volume":"43","author":"R Engbert","year":"2003","unstructured":"Engbert R, Kliegl R (2003) Microsaccades uncover the orientation of covert attention. Vision Res 43:1035\u20131045","journal-title":"Vision Res"},{"key":"7320_CR13","doi-asserted-by":"publisher","first-page":"16","DOI":"10.1016\/S1364-6613(00)01568-0","volume":"5","author":"AK Engel","year":"2001","unstructured":"Engel AK, Singer W (2001) Temporal binding and the neural correlates of sensory awareness. Trends Cogn Sci 5:16\u201325","journal-title":"Trends Cogn Sci"},{"issue":"3","key":"7320_CR14","doi-asserted-by":"publisher","first-page":"18","DOI":"10.1167\/18.3.18","volume":"18","author":"Y Fang","year":"2018","unstructured":"Fang Y, Gill C, Poletti M, Rucci M (2018) Monocular microsaccades: do they really occur? J Vision 18(3):18. https:\/\/doi.org\/10.1167\/18.3.18","journal-title":"J Vision"},{"issue":"6","key":"7320_CR15","doi-asserted-by":"publisher","first-page":"e0310619","DOI":"10.1371\/journal.pone.0310619","volume":"20","author":"L Friedman","year":"2025","unstructured":"Friedman L, Komogortsev OV (2025) Fixation drift increases as a function of time-on-task in a brief saccade tracking study. PLoS ONE 20(6):e0310619. https:\/\/doi.org\/10.1371\/journal.pone.0310619","journal-title":"PLoS ONE"},{"key":"7320_CR16","doi-asserted-by":"publisher","first-page":"209","DOI":"10.1146\/annurev.neuro.051508.135603","volume":"32","author":"P Fries","year":"2009","unstructured":"Fries P (2009) Neuronal gamma-band synchronization as a fundamental process in cortical computation. Annu Rev Neurosci 32:209\u2013224","journal-title":"Annu Rev Neurosci"},{"key":"7320_CR17","doi-asserted-by":"publisher","first-page":"1483","DOI":"10.1097\/01.wnr.0000175611.26485.c8","volume":"16","author":"K Heinen","year":"2005","unstructured":"Heinen K, Jolij J, Lamme VAF (2005) Figure-ground segregation requires two distinct periods of activity in V1: a transcranial magnetic stimulation study. NeuroReport 16:1483\u20131487","journal-title":"NeuroReport"},{"key":"7320_CR18","doi-asserted-by":"publisher","first-page":"12958","DOI":"10.1038\/s41598-017-13489-8","volume":"7","author":"CJJ Herrmann","year":"2017","unstructured":"Herrmann CJJ, Metzler R, Engbert R (2017) A self-avoiding walk with neural delays as a model of fixational eye movements. Sci Rep 7:12958. https:\/\/doi.org\/10.1038\/s41598-017-13489-8","journal-title":"Sci Rep"},{"issue":"11","key":"7320_CR19","doi-asserted-by":"publisher","first-page":"1335","DOI":"10.1167\/jov.20.11.1335","volume":"20","author":"J Intoy","year":"2020","unstructured":"Intoy J, Victor JD, Rucci M (2020) Active task-dependent control of ocular drift during natural fixation. J vis 20(11):1335. https:\/\/doi.org\/10.1167\/jov.20.11.1335","journal-title":"J vis"},{"key":"7320_CR20","first-page":"5792","volume":"59","author":"D Ivanchenko","year":"2018","unstructured":"Ivanchenko D, Hafed Z, Schaeffel D (2018) How correlated are drifts in both eyes during fixational eye movements? Invest Ophthalmol vis Sci 59:5792","journal-title":"Invest Ophthalmol vis Sci"},{"issue":"13","key":"7320_CR21","doi-asserted-by":"publisher","first-page":"e1815232024","DOI":"10.1523\/JNEUROSCI.1815-23.2024","volume":"44","author":"F Khademi","year":"2024","unstructured":"Khademi F, Zhang T, Baumann MP, Malevich T, Yu Y, Hafed ZM (2024) Visual feature tuning properties of short-latency stimulus-driven ocular position drift responses during gaze fixation. J Neurosci 44(13):e1815232024. https:\/\/doi.org\/10.1523\/JNEUROSCI.1815-23.2024","journal-title":"J Neurosci"},{"key":"7320_CR22","doi-asserted-by":"publisher","first-page":"1549","DOI":"10.1038\/nn.2663","volume":"13","author":"HK Ko","year":"2010","unstructured":"Ko HK, Poletti M, Rucci M (2010) Microsaccades precisely relocate gaze in a high visual acuity task. Nat Neurosci 13:1549\u20131553. https:\/\/doi.org\/10.1038\/nn.2663","journal-title":"Nat Neurosci"},{"key":"7320_CR23","doi-asserted-by":"publisher","first-page":"510","DOI":"10.1016\/j.cub.2012.01.050","volume":"22","author":"X Kuang","year":"2012","unstructured":"Kuang X, Poletti M, Victor JD, Rucci M (2012) Temporal encoding of spatial information during active visual fixation. Curr Biol 22:510\u2013514","journal-title":"Curr Biol"},{"key":"7320_CR24","doi-asserted-by":"publisher","first-page":"571","DOI":"10.1016\/S0166-2236(00)01657-X","volume":"23","author":"VAF Lamme","year":"2000","unstructured":"Lamme VAF, Roelfsema PR (2000) The distinct modes of vision offered by feedforward and recurrent processing. Trends Neurosci 23:571\u2013579","journal-title":"Trends Neurosci"},{"key":"7320_CR25","doi-asserted-by":"publisher","first-page":"797","DOI":"10.1016\/0042-6989(92)90022-B","volume":"32","author":"VA Lamme","year":"1992","unstructured":"Lamme VA, Van Dijk BW, Spekreijse H (1992) Texture segregation is processed by primary visual cortex in manand monkey. Evid VEP Exp Vision Res 32:797\u2013807","journal-title":"Evid VEP Exp Vision Res"},{"issue":"5","key":"7320_CR26","doi-asserted-by":"publisher","first-page":"e2004132","DOI":"10.1371\/journal.pbio.2004132","volume":"16","author":"E Lowet","year":"2018","unstructured":"Lowet E, Gips B, Roberts MJ, De Weerd P, Jensen O, van der Eerden J (2018) Microsaccade-rhythmic modulation of neural synchronization and coding within and across cortical areas V1 and V2. PLoS Biol 16(5):e2004132. https:\/\/doi.org\/10.1371\/journal.pbio.2004132","journal-title":"PLoS Biol"},{"key":"7320_CR27","doi-asserted-by":"publisher","first-page":"e57595","DOI":"10.7554\/eLife.57595","volume":"9","author":"T Malevich","year":"2020","unstructured":"Malevich T, Buonocore A, Hafed ZM (2020) Rapid stimulus-driven modulation of slow ocular position drifts. Elife 9:e57595. https:\/\/doi.org\/10.7554\/eLife.57595","journal-title":"Elife"},{"issue":"6","key":"7320_CR28","doi-asserted-by":"publisher","first-page":"24086","DOI":"10.1038\/srep24086","volume":"11","author":"T Masquelier","year":"2016","unstructured":"Masquelier T, Portelli G, Kornprobst P (2016) Microsaccades enable efficient synchrony-based coding in the retina: a simulation study. Sci Rep 11(6):24086. https:\/\/doi.org\/10.1038\/srep24086","journal-title":"Sci Rep"},{"issue":"6","key":"7320_CR29","doi-asserted-by":"publisher","first-page":"e0128428","DOI":"10.1371\/journal.pone.0128428","volume":"10","author":"MB McCamy","year":"2015","unstructured":"McCamy MB, Otero-Millan J, Leigh RJ, King SA, Schneider RM, Macknik SL, Martinez-Conde S (2015) Simultaneous recordings of human microsaccades and drifts with a contemporary video eye tracker and the search coil technique. PLoS ONE 10(6):e0128428. https:\/\/doi.org\/10.1371\/journal.pone.0128428","journal-title":"PLoS ONE"},{"key":"7320_CR30","doi-asserted-by":"publisher","first-page":"1173","DOI":"10.1016\/j.visres.2010.04.007","volume":"50","author":"A Pastukhov","year":"2010","unstructured":"Pastukhov A, Braun J (2010) Rare but precious: microsaccades are highly informative about attentionalal location. Vision Res 50:1173\u20131184. https:\/\/doi.org\/10.1016\/j.visres.2010.04.007","journal-title":"Vision Res"},{"key":"7320_CR31","doi-asserted-by":"publisher","first-page":"214","DOI":"10.3758\/s13428-025-02731-1","volume":"57","author":"AR Pijpaert","year":"2025","unstructured":"Pijpaert AR, Goossens HHLMJ, van Dijk BW et al (2025) A validation study on the accuracy and precision of gaze and vergence using stereoscopic eye-tracking technology. Behav Res 57:214. https:\/\/doi.org\/10.3758\/s13428-025-02731-1","journal-title":"Behav Res"},{"issue":"24","key":"7320_CR32","doi-asserted-by":"publisher","first-page":"6086","DOI":"10.1016\/j.cub.2025.11.007","volume":"35","author":"KS Prahalad","year":"2025","unstructured":"Prahalad KS, Clark AM, Moon B, Roorda A, Tiruveedhula P, Harmening W, Gutnikov A, Jenks SK, Kapisthalam S, Rucci M, Rolland JP, Poletti M (2025) Non-uniform signal pooling across the foveola. Curr Biol 35(24):6086-6099.e4. https:\/\/doi.org\/10.1016\/j.cub.2025.11.007","journal-title":"Curr Biol"},{"key":"7320_CR33","doi-asserted-by":"publisher","first-page":"852","DOI":"10.1167\/3.11.18","volume":"3","author":"M Rucci","year":"2003","unstructured":"Rucci M, Desbordes G (2003) Contributions of fixational eye movements to the discrimination of briefly presented stimuli. J Vis 3:852\u2013864","journal-title":"J Vis"},{"issue":"4","key":"7320_CR34","doi-asserted-by":"publisher","first-page":"195","DOI":"10.1016\/j.tins.2015.01.005","volume":"38","author":"M Rucci","year":"2015","unstructured":"Rucci M, Victor JD (2015) The unsteady eye: an information-processing stage, not a bug. Trends Neurosci 38(4):195\u2013206. https:\/\/doi.org\/10.1016\/j.tins.2015.01.005","journal-title":"Trends Neurosci"},{"key":"7320_CR35","doi-asserted-by":"publisher","first-page":"49","DOI":"10.1016\/S0896-6273(00)80821-1","volume":"24","author":"W Singer","year":"1999","unstructured":"Singer W (1999) Neuronal synchrony: a versatile code of the definition of relations? Neuron 24:49\u201365","journal-title":"Neuron"},{"key":"7320_CR36","doi-asserted-by":"publisher","first-page":"103138","DOI":"10.1016\/j.concog.2021.103138","volume":"92","author":"M Sol\u00e9 Puig","year":"2021","unstructured":"Sol\u00e9 Puig M, Romeo A, Sup\u00e8r H (2021) Vergence eye movements during figure-ground perception. Conscious Cogn 92:103138","journal-title":"Conscious Cogn"},{"key":"7320_CR37","doi-asserted-by":"publisher","first-page":"304","DOI":"10.1038\/85170","volume":"4","author":"H Sup\u00e8r","year":"2001","unstructured":"Sup\u00e8r H, Spekreijse H, Lamme VAF (2001) Two distinct modes of sensory processing observed in monkey primary visual cortex (V1). Nat Neurosci 4:304\u2013310","journal-title":"Nat Neurosci"},{"key":"7320_CR38","doi-asserted-by":"publisher","first-page":"3407","DOI":"10.1523\/JNEUROSCI.23-08-03407.2003","volume":"23","author":"H Sup\u00e8r","year":"2003","unstructured":"Sup\u00e8r H, van der Togt C, Spekreijse H, Lamme VAF (2003) Internal state of monkey primary visual cortex (V1) predicts figure-ground perception. J Neurosci 23:3407\u20133414","journal-title":"J Neurosci"},{"key":"7320_CR39","doi-asserted-by":"publisher","first-page":"5461","DOI":"10.1523\/JNEUROSCI.20-14-05461.2000","volume":"20","author":"WM Usrey","year":"2000","unstructured":"Usrey WM, Alonso JM, Reid RC (2000) Synaptic interactions between thalamic inputs to simple cells in cat visual cortex. J Neurosci 20:5461\u20135467","journal-title":"J Neurosci"},{"key":"7320_CR40","doi-asserted-by":"publisher","first-page":"136","DOI":"10.1093\/cercor\/bhi093","volume":"16","author":"C Van der Togt","year":"2006","unstructured":"Van der Togt C, Kalitzin S, Spekreijse H, Lamme VAF, Sup\u00e8r H (2006) Synchrony dynamics in monkey V1 predict success in visual detection. Cereb Cortex 16:136\u2013148","journal-title":"Cereb Cortex"}],"container-title":["Experimental Brain Research"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s00221-026-07320-y.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1007\/s00221-026-07320-y","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s00221-026-07320-y.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,14]],"date-time":"2026-08-14T05:21:45Z","timestamp":1786684905000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1007\/s00221-026-07320-y"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,7,23]]},"references-count":38,"journal-issue":{"issue":"8","published-print":{"date-parts":[[2026,8]]}},"alternative-id":["7320"],"URL":"https:\/\/doi.org\/10.1007\/s00221-026-07320-y","relation":{"has-preprint":[{"id-type":"doi","id":"10.21203\/rs.3.rs-8699876\/v1","asserted-by":"object"}]},"ISSN":["0014-4819","1432-1106"],"issn-type":[{"value":"0014-4819","type":"print"},{"value":"1432-1106","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,7,23]]},"assertion":[{"value":"26 January 2026","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"14 May 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"23 July 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"HS is co-founder of Braingaze.","order":1,"name":"Ethics","label":"Competing interest","group":{"name":"EthicsHeading","label":"Declarations"}}],"article-number":"162"}}